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Harikrishnan_Vasudevan_Calotes_andamanensis_HERPETOZOA.qxd 21.06.2013 12:18 Seite 1

herpeToZoA 26 (1/2): 3 - 13 3 wien, 30. Juni 2013

rediscovery of andamanensis Boulenger , 1891, and assessment of its generic allocation (: sauria: )

wiederentdeckung von Calotes andamanensis Boulenger , 1891 und die Beurteilung seiner gattungszugehörigkeit (squamata: sauria: Agamidae)

surenDrAn hArIKrIshnAn & KArThIKeyAn VAsuDeVAn

KurZFAssung

Die nur vom holotypus bekannte draconine Agame Calotes andamanensis Boulenger , 1891, wurde auf den Andamanen () wiedergefunden. Die Autoren legen morphometrische und meristische Daten von sechs exemplaren von drei Inseln des Andamanen-Archipels vor und machen ergänzende Angaben zur Beschreibung des Typusexemplares. nach der äußeren Morphologie der echsen ist ihre Zuordnung zur gattung Calotes nicht zutr - effend, vielmehr sollten sie in die gattung gestellt werden. ABsTrACT

The little known draconine agamid Calotes andamanensis Boulenger , 1891, was rediscovered from the Andaman Islands (India). The authors present morphometric and meristic data from six specimens originating from three islands of the Andaman Archipelago, and thereby add information to the description of the type speci - men. The lizard’s external morphology suggests that its generic assignment to Calotes in its present definition can - not be maintained and that this should better be included in the Pseudocalotes . Key worDs reptilia: squamata: sauria: Agamidae; Calotes , Pseudocalotes andamanensis , lizard, systematics, taxo- nomy, Andaman Islands, nicobar Islands, India, Indo-

InTroDuCTIon

The enigmatic agamid lizard Calotes in his “ liocephalus group”, which otherwise andamanensis (Boulenger , 1891 ) was de- included sri lankan endemic species. This scribed from a single specimen in the col - species was never again reported from the lection of the Zoologisk Museum, uni- Andaman and nicobar Islands or elsewhere. versity of Copenhagen (ZMuC) in Den - IshwAr & D As (1999) assigned a mark. The specimen was originally collect - specimen of Calotes from the Kalakkad- ed by Frederick Adolph De roepsTorFF , Mundanthurai Tiger reserve (KMTr) in the officer-in-charge of the British camp at southern part of the western ghats in India nancowry harbor, Andaman and nicobar to C. andamanensis , and suggested that the Islands ( sMITh 1935; K rIshnAn 2008). type locality “Andaman Islands” was prob - Boulenger (1891) did not elaborate on the ably assigned in error. Consequently, it was affinities of this species, but considered the not listed in a subsequent list of Andaman species to be closely related to the sri Island herpetofauna (D As 1999). however, lankan species Boulen- the specimen from KMTr was later de- ger , 1885 , and listed the differences be - scribed as Calotes aurantolabium KrIsh- tween C. andamanensis and C. liolepis viz., nAn , 2008. In this publication, KrIshnAn much larger scales and the presence of a (2008) discussed the differences of C. anda - pair of spine-like scales on each side of the manensis to other south Asian members of back of the head in the latter (B oulenger , the genus Calotes and provided a refined 1891). s MITh (1935) followed Boulenger ’s description of its holotype. Moreover, he (1891) opinion and placed C. andamanensis pointed out that the collector of the type Harikrishnan_Vasudevan_Calotes_andamanensis_HERPETOZOA.qxd 21.06.2013 12:18 Seite 2

4 s. hArIKrIshnAn & K. V AsuDeVAn

specimen of C. andamanensis , De roeps- tified as C. andamanensis , but added that it TorFF , was posted in the nicobar Islands, is in a bad state of preservation and cannot thus suggesting that the specimen probably be diagnosed. This specimen was collected originated from there and not the Andaman from “Middle Andaman by Dr. gAnApAThy Islands. Although there is a submerged in 1964” ( VArAD gIrI , pers. comm.). mountain ridge running from the Andamans here, the authors report on six draco - to the nicobars, i. e., Arakan yomas to nine from the Andaman Islands, pro - Mentawei Islands, these two island groups vide morphometric and meristic data on are separated by a deep channel known as these specimens and compare it with de- the Ten Degree Channel, which acts as a scriptions by Boulenger (1891) and KrIsh- zoogeographical barrier ( sMITh 1940; D As nAn (2008), and also with other and espe - 1999). KrIshnAn (2008) also mentioned a cially closely related species, as well as specimen in the collection of the Bombay Pseudocalotes species. natural history society (BnhM 710), iden -

MATerIAls AnD MeThoDs

The lizards, stored at the wildife Insti - 0.01 mm, the long distances snout-vent tute of India, Vertebrate Collections at Dehra length (sVl), leg length (ll), torso length Dun, uttarakhand, India (wII), originated (Trl), torso width (Trw), torso height (Trh) from three islands of the Andaman group: and tail length (Tl) to the nearest 0.1 mm. wII 001 – long Island (Middle Anda - The following measurements were record - man Islands), adult male, collected on 9 ed: snout-vent length (sVl - from tip of April, 2010, by harikrishnan, s. & Chand- snout to anterior edge of cloacal opening); ramouli, s. r. tail length (Tl - from posterior edge of cloa - wII 126 – rutland, adult female, col - cal opening to tip of tail); head length (hl - lected on 21 April, 2011, by harikrishnan, from tip of snout to posterior border of tym - s. & Chandramouli, s. r. panum, following hAllerMAnn [2005] as wII 055 – south Andaman, road to this avoids the potential ambiguity when Mt. harriet national park, near hopetown, head length is measured from snout tip to adult male, collected on 1 March, 2011, by occiput); head width (hw - maximum harikrishnan, s. & Chandramouli, s. r. width of head behind orbit); head depth (hD wII 056 – south Andaman, road to - height of head behind orbit), snout length Mt. harriet national park, near hopetown, (sl - distance between tip of snout and ante - adult male, collected on 1 March, 2011, by rior edge of orbit); jaw length (Jl - distance harikrishnan, s. & Chandramouli, s. r. from tip of lower jaw to the posterior mar - wII 072 – south Andaman, hopetown, gin of lower jaw); orbit diameter (oD - dis - adult female, collected on 8 March, 2011, tance between anterior and posterior edges by harikrishnan, s. of orbit); tympanum diameter (TD - dis - wII 125 – south Andaman, hopetown, tance between anterior and posterior edges adult male, collected on 20 March, 2011, by of tympanum); torso length (Trl - distance harikrishnan, s. between axilla and groin); torso width (Trw The long Island specimen was col - - width of the body at the middle of Trl); lected from a tree trunk at a height of two torso height (Trh - height of the body at the meters, the rutland specimen was found on middle of Trl); upper arm length (hum - a tree at four meters above ground. Two of distance between shoulder joint and elbow); the specimens from south Andaman were forearm length (rad - distance between captured on the ground, one on the trunk of elbow and wrist); length of fingers exclud - a tree three meters above the ground and ing the claw (F1, F2, F3, F4 & F5 numbered one climbing on the outer wall of a building successively); thigh length (Fem - distance (2.2 m above ground). between the groin and the knee); shank Measurements were taken with Mitu - length (Tib - distance between knee and toyo ® dial vernier calipers to the nearest ankle); length of toes excluding the claw Harikrishnan_Vasudevan_Calotes_andamanensis_HERPETOZOA.qxd 21.06.2013 12:18 Seite 3

rediscovery of Calotes andamanensis Boulenger , 1891 5

(T1, T2, T3, T4 & T5 numbered successive - different. specimen numbers given are the ly); leg length (ll - distance between the field catalogue numbers in the collection of groin and the tip of the fourth toe, excluding the wildlife Institute of India, Dehra Dun. the claw; measured by stretching the limbs). These specimens are part of an ongoing scales around the body were counted at half study and will be deposited in the Zoo - the distance between fore and hind limbs logical survey of India, Kolkata (ZsI) col - (sMITh , 1935). Canthals were counted along lection on completion of the study. Another the canthus rostralis between nasal and first specimen, a male found in wandoor (south supraorbital at the anterior edge of orbit. Andaman Island) in January 2009, was col - only the scales in contact with the mental lected and deposited in the collection of the were regarded as postmentals, excluding the Zoological survey of India, Marine region- first infralabials. al station, port Blair, Anda man and nicobar In the following description of six Islands, India (ZsI 10641). This was the freshly collected specimens, the character first specimen of this species collected by the states of the holotype of C. andamanensis authors of this paper. subsequent efforts to are indicated in parentheses, if they were locate this specimen were unsuccessful.

resulTs

Description of the draconine lizard sample 57-62 (67 in the holotype) longitudi - (Figs. 1A, 1B, 2A, 2B) nal scale rows around midbody; scales of the upper 4-7 rows directed posterodorsally, head elongate (hw:hl = 0.59); pos - remaining lateral scales directed postero- terior part of jaw region swollen, particular - ventrally; scales of uppermost paravertebral ly in males; nostril placed within enlarged rows enlarged, heterogeneous in shape, pos - nasal; rostral in contact with 4-6 scales on terior ones feebly keeled; the other lateral snout excluding the first supralabial on scales smaller; dorsal and lateral scales either side; 4-5 canthals between nasal and smooth except above pelvis; ventrals strong- first supraciliary; 7-8 supraciliaries; 3-4 ly keeled, slightly irregular, smaller than dor - interorbitals; upper head scales obtusely sals but of similar size as laterals; dorsal tail keeled; upper snout scales flattened except scales weakly keeled; ventral tail scales the median row which is keeled, becoming strongly keeled; nuchal crest well developed, progressively keeled towards posterior part with 11-14 spines on a slightly raised flap of of head; rostral in contact with 3 enlarged skin (15 in holotype); dorsal crest a mere ser - scales on the tip of the snout; these scales rated ridge (in the holotype described as ‘a are separated from the nasal by one scale; 3- serrated ridge’ [ Boulenger 1891] or ‘weak - 5 enlarged scales on occiput, their degree of ly developed’ [ KrIshnAn 2008]); a shallow enlargement highly variable; tympanum antehumeral pit and a weak antehumeral exposed, its dia meter less than half the orbit fold, not extending across the throat. diameter (TD: oD = 0.39); 3-5 enlarged number of enlarged keeled scales on scales between orbit and tympanum; scales posterior part of thigh 0-6, scales of this below this also slightly enlarged; 4-5 inter - region being of irregular size; 3rd and 4th nasals; 9-10 supralabials; nasal in broad fingers of similar length; fourth toe longest; contact with supralabials except in wII 001, 20-23 lamellae underneath third finger; 27- in which the nasal is separated from supral - 30 lamellae underneath fourth toe; first pha - abials by a row of small scales; one row of lange of third toe with enlarged triangular enlarged scales between supralabials and lamellae (Fig. 3); hind limb relatively long orbit, with one or two rows of smaller scales (70-75 % of sVl); tail relatively long (238- above and below; 10-12 infralabials; 2 post - 265 % of sVl), feebly compressed at the mentals except in wII 001 which has 3; base; a gular pouch present in males, reach - postmentals followed by a row of enlarged ing the chest (Fig. 4B). scales separated from the infralabials by a Coloration in life (Figs. 4A, 4B).- row of slightly smaller scales. Body pale green above and yellowish Harikrishnan_Vasudevan_Calotes_andamanensis_HERPETOZOA.qxd 21.06.2013 12:18 Seite 4

6 s. hArIKrIshnAn & K. V AsuDeVAn Harikrishnan_Vasudevan_Calotes_andamanensis_HERPETOZOA.qxd 21.06.2013 12:18 Seite 5

rediscovery of Calotes andamanensis Boulenger , 1891 7

Table 1: Metric data (mm) from six specimens of Pseudocalotes andamanensis (Boulenger , 1891) from the Andaman Islands studied in this paper. Data of the holotype (ZMuC 36944) was taken from KrIshnAn (2008). For definition of measured parameters see Materials and Methods. M – Male, F – Female. Tab. 1: Metrische Daten (mm) von sechs in dieser Arbeit untersuchten Pseudocalotes andamanensis (Boulenger , 1891) von den Andamanen Inseln. Die Daten des holotypus (ZMuC 36944) stammen aus KrIshnAn (2008). Die Meßstreckendefinitionen sind in Material und Methoden angegeben. M – Männchen, F – weibchen.

parameter wII 001 wII 055 wII 056 wII 072 wII 125 wII 126 ZMuC 36944 sex MM M F M F M sVl 84.7 86.3 86.9 76.2 83.6 72.4 84.1 Tl 207.2 205.4 211.0 220.1 204.2 173.1 - Tw 7.05 7.96 7.20 6.96 7.55 6.00 - hl 24.23 23.68 23.39 21.55 23.62 19.76 23.69 hw 13.82 13.76 14.14 12.87 13.54 11.68 13.62 hD 13.66 12.81 10.92 12.05 13.10 11.40 12.42 sl 10.44 9.96 10.58 9.24 10.12 8.61 - Jl 25.93 25.47 25.55 23.13 26.99 23.05 - oD 7.39 7.74 6.66 7.23 7.04 6.30 7.39 TD 3.16 2.85 2.48 2.70 3.03 2.48 2.70 Trl 39.4 39.0 38.1 35.2 38.0 33.1 44.42 Trw 12.6 13.9 11.1 14.9 15.0 17.1 - Trh 15.0 15.5 16.3 16.6 15.3 16.5 - hum 13.83 13.65 14.75 16.74 18.27 15.00 15.44 rad 14.66 15.56 15.4 15.53 16.21 12.71 15.33 F1 2.78 3.04 3.43 3.85 3.71 3.06 - F2 5.28 5.68 6.34 6.32 5.80 4.43 - F3 8.42 8.87 9.45 9.93 8.42 7.05 - F4 8.38 9.04 9.29 9.97 9.11 7.05 - F5 4.49 5.26 6.49 5.74 5.33 4.33 - Fem 19.81 19.05 21.03 19.37 20.54 14.34 19.28 Tib 19.77 19.60 21.16 20.90 21.31 16.33 19.96 T1 2.92 3.58 3.28 4.20 3.58 2.73 - T2 6.15 6.64 6.03 6.20 5.53 4.62 - T3 10.32 10.98 11.35 11.59 11.29 9.88 - T4 13.75 13.84 13.65 14.12 13.10 11.54 - T5 8.71 9.29 9.63 10.38 9.11 7.68 - ll 62.3 60.1 64.1 54.0 62.4 54.3 -

below; upper lip white, with a white stripe fighting each other had the inner parts of extending to behind tympanum; body with lips and tongue colored bright yellow. All white spots and vertical bars; tail banded specimens turned pale bluish-white or with pale green and white; the green color brown in alcohol. rapidly disappears when stressed and the This species appears to be arboreal, whole turns brown on capture. preferring the crown or canopy of trees. All gular pouch white or yellow in males, individuals were sighted during the relative - absent in females. Two of the males found ly drier season between January and April.

______Figs. 1 - 2 (opposite page) / Abb. 1 - 2 (gegenüberliegende seite).

Fig. 1: Pseudocalotes andamanensis (Boulenger , 1891) – lateral view of head. A - ZMuC 36944, holotype; B - wII 055. Abb. 1: Pseudocalotes andamanensis (Boulenger , 1891) – Kopf von lateral. A - ZMuC 36944, holotypus; B - wII 055.

Fig. 2: Pseudocalotes andamanensis (Boulenger , 1891) – Dorsal view of head. A - ZMuC 36944, holotype; B - wII 055. Abb. 2: Pseudocalotes andamanensis (Boulenger , 1891) – Dorsalansicht des Kopfes. A - ZMuC 36944, holotypus; B - wII 055. Harikrishnan_Vasudevan_Calotes_andamanensis_HERPETOZOA.qxd 21.06.2013 12:18 Seite 6

8 s. hArIKrIshnAn & K. V AsuDeVAn

thus, speculated that the type probably orig - inated in the nicobar and not in the Andaman Islands. This is however, contra - dicted by De roepsTorFF himself who reported his interactions with people of the tribe ‘great Andamanese’ near port Blair, south Andaman Island (De roepsTorFF 1875) . Although De roepsTorFF did not elaborate on the natural history of these islands, the authors believe that mentioning the above interaction is sufficient evidence for De roepsTorFF having visited the Andaman Islands. Also, apart from the type of C. andamanensis , there exists a number of other herpetological specimens in the col - lection of the Zoologisk Museum, university of Copenhagen, collected by De roepsTorFF , and labeled as being from the Andaman Islands. These include Laticauda colubrina (s ChneIDer , 1799) , Ophiophagus hannah (C AnTor , 1836) , Naja sagittifera wAll , 1913 (labeled Naja kaouthia lesson , 1831 , for synonymy comp. wüsTer [1998]), Bungarus andamanensis BIswAs & sAnyAl , 1978 (locality data says “Anda- man, south”, presumably south Andaman Island), Trimeresurus andersonii Theo - BAlD , 1868 (labeled Trimeresurus purpureo- Fig. 3: Pseudocalotes andamanensis (Boulenger , maculatus (g rAy , 1832) , of which the 1891) – enlarged and modified lamellae on the Andaman taxon was previously considered leading edge of the third toe (wII 126). a , comp. MAlhoTrA & T horpe Abb. 3: Pseudocalotes andamanensis (Boulenger , [1997]), Dendrelaphis pictus (Andaman 1891) – Vergrößerte umgestaltete lamellen an der unterkante der dritten Zehe (wII 126). Islands population now referred to as Dendrelaphis andamanensis (AnDerson , 1871 ) [ Vogel & VAn rooIJen 2011]). The above material is clear evidence that De Apart from the specimens mentioned here, roepsTorFF did indeed procure specimens eight more individuals of this type were collected in the Andaman Islands. There- observed, all on high branches of trees, up fore, the type locality, given as “Andaman to 30 m above ground. on the ground, they Islands” is regarded to be correct by the moved clumsily and seemed to be relatively authors of this paper. slow moving in the trees also. nothing else is known about the ecology of this species. generic allocation of Boulenger ’s (1891) Calotes andamanensis Comments on the type locality of Calotes andamanensis Describing Calotes andamanensis , Boulenger (1891) mentioned that it most According to the original description closely resembled Calotes liolepis günT- (Boulenger 1891), the holotype of C. anda - her , 1872, a species found in , but manensis was collected on the Andaman without specifying why he thought so. Islands by Frederick Adolph De roep sTorFF . sMITh (1935) followed Boulenger ’s opin - KrIshnAn (2008) stated that De roepsTorFF ion and placed C. andamanensis in his ‘ lio - was posted in the nicobar Islands and prob - cephalus group’. This concept of Calotes ably never visited the Andaman Islands and, has changed since, and members of these Harikrishnan_Vasudevan_Calotes_andamanensis_HERPETOZOA.qxd 21.06.2013 12:18 Seite 7

rediscovery of Calotes andamanensis Boulenger , 1891 9

Table 2: Meristic data from six specimens of Pseudocalotes andamanensis (Boulenger , 1891) from the Andaman Islands compared with the holotype (ZMuC 36944). Data of the holotype was taken from Boulenger (1891) and KrIshnAn (2008). For definition of parameters see Materials and Methods. M – Male, F – Female. The counts on left and right sides of body are separated by a comma. Table 2: Meristische Daten von sechs in dieser Arbeit untersuchten Pseudocalotes andamanensis (Boulenger , 1891) von den Andamanen sowie die des holotypus (ZMuC 36944) im Vergleich. Daten des holotypus aus Boulenger (1891) and KrIshnAn (2008). Die Meßstreckendefinitionen sind in Material und Methoden angegeben. M – Männchen, F – weibchen. rechte und linke Zählwerte sind durch Kommas getrennt.

parameter wII 001 wII 055 wII 056 wII 072 wII 125 wII 126 ZMuC 36944 sex MMMFM F M supralabials 10, 10 10, 10 9, 9 10,10 10, 10 10,10 10, 10 Infralabials 10, 11 10, 10 9, 9 12, 11 12, 10 10, 11 10, 13 Internasals 4 5555 5 - Canthals 4 5, 4 5, 4 55 5 - supraciliaries 8778,7 7, 8 6, 8 - Interorbitals 44444 3 4 postmentals 32222 2 2 upper head scales Keeled Keeled Keeled Keeled Keeled Keeled Keeled upper snout scales smooth smooth smooth smooth smooth smooth - enlarged scales 34445 4 5 on temple scales around 62 58 60 57 57 57 67 *) midbody nuchal spines 13 14 13 13 11 12 15 lamellae under 20 22 20 22 23 22 22 3rd finger lamellae under 29 29 27 29 29 30 28 4th toe enlarged keeled 0135/6 423 scales on thigh

*) The original scale count given by Boulenger (1891) was 63. however, sMITh (1943) and KrIshnAn (2008) reported 67.

groups are now assigned to the genera nuchal, dorsal and caudal crests and en- KAup , 1827, Pseudocalotes larged keeled dorsal and lateral scales FITZInger , 1843 and Calotes CuVIer , 1817 (sMITh 1935; M Ahony 2010). The absence (MooDy 1980; h AllerMAnn & B öhMe of a longitudinal skin fold extending from 2000; M Ahony 2010). In their review of the angle of the jaw to the shoulder, ventrals Pseudocalotes , hAllerMAnn & B öhMe that are smaller than dorsals and the rela - (2000) pointed out morphological charac - tively short tail clearly indicates that anda - ters to separate the genera Pseudocalotes , manensis is not a member of the genus Calotes and Bronchocela . If this classifica - Bronchocela as defined by hAllerMAnn & tion is accepted, Calotes andamanensis is BöhMe (2000) and hAllerMAnn (2005) . clearly not a member of the genus Calotes In showing a row of enlarged scales CuVIer , 1817 due to the lack of postorbital between orbit and tympanum, heteroge - and supratympanic spines, and the presence neous dorsal scales, enlarged triangular of a heterogenous body scalation, a relative - lamellae under the third toe and lacking an ly long and narrow head and a row of enlarged tail base in males, this species is enlarged scales between orbit and supralabi - most similar to species of the genus Pseudo - als. MAhony (2010) discussed the impor - calotes , which are widespread in southeast tance of certain morphological characters in Asia ( MooDy 1980; h AllerMAnn & B öh- the systematics of draconine agamids and Me 2000; h AllerMAnn & M CguIre 2001; pointed out that the subocular row of hAllerMAnn et al. 2009; MAhony 2010). enlarged scales is characteristic of several In having relatively slender and long limbs, insular genera as well as Salea grAy , 1845 . the present specimens of the Andaman Andamanensis is certainly not a member of Islands are most similar to Pseudocalotes the genus Salea as it lacks its conspicuous tympanistriga (g rAy , 1831) from Java and Harikrishnan_Vasudevan_Calotes_andamanensis_HERPETOZOA.qxd 21.06.2013 12:18 Seite 8

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Fig. 4: Pseudocalotes andamanensis (Boulenger , 1891) males in life. A - wII 055; B - wII 056. Abb. 4: Pseudocalotes andamanensis (Boulenger , 1891). Männchen. A - wII 055; B - wII 056. Harikrishnan_Vasudevan_Calotes_andamanensis_HERPETOZOA.qxd 21.06.2013 12:18 Seite 9

rediscovery of Calotes andamanensis Boulenger , 1891 11

sumatra, whereas it appears closest to P. Diagnosis.- relatively long head (hw: microlepis (B oulenger , 1888) from the hl = 0.59); 56-67 longitudinal scale rows Indo-Chinese peninsula and P. larutensis around midbody; dorsals and laterals hAllerMAnn & M CguIre , 2001 from west - smooth, sometimes weakly keeled near the ern in mid-body scale counts and sacral region; ventrals strongly keeled; dor - scalation ( hAllerMAnn & M CguIre 2001 ). sals of 4-7 paravertebral longitudinal rows From P. tympanistriga , P. andamanensis larger than laterals, of irregular shape, differs in having a wider head (hw/hl 0.59 pointing posterodorsally; laterals pointing in P. andamanensis , 0.52 in P. tympanistri - posteroventrally; laterals and ventrals of ga ), longer hind limbs (70-75 % of sVl in similar size; ventrals slightly irregular; a P. andamanensis , 68 % in P. tympanistriga ), row of enlarged scales between supralabials 9-10 supralabials and 9-12 infralabials (10- and orbit, bordered by one or two smaller 13 supralabials and 9-11 infralabials in P. scale rows; gular scales smaller than ven - tympanistriga ). In having enlarged triangu - trals, weakly keeled; gular pouch present in lar lamellae on the leading edge of the third males; antehumeral fold/pit weakly devel - toe, this species is similar to the Indo- oped; nuchal crest composed of 11-15 Chinese species group of Pseudocalotes . lanceolate spines; dorsal crest a denticulate It should be remembered that most ridge; enlarged conical lamellae under the draconine agamid genera are defined solely leading edge of third toe; 27- 30 lamellae based on phenetic characters. In several under fourth toe; hind limb length 70-75 % traits, the species from the Andamans differs of sVl; tail length 238-265 % of sVl, from other Pseudocalotes , most notably its slightly compressed at the base. proportionately longer limbs. only P. tym - panistriga is known to have similar limb Distribution, habits and habitat proportions. In addition, most species of Pseudocalotes , except P. saravacensis Inger Pseudocalotes andamanensis (Bou - & s TueBIng , 1994, are inhabitants of moun - lenger , 1891) is an arboreal species, usual - tain forests, whereas the Andamanese spe- ly found on the trunk of trees near the cies is known from sea level to about 200 m crown, or on branches. This species is a.s.l. Moreover, it exhibits a weak ante - known from south Andaman Island, humeral fold which is otherwise present in rutland Island and long Island. owing to genera such as Calotes CuVIer , 1817 (some its arboreal habits, it is rarely sighted and is species), Psammophilus FITZInger , 1843, likely to occur in more islands in the KAup , 1825, Japalura grAy , Andaman Islands. The only other native 1853, Acanthosaura grAy , 1831, etc. how - agamid species that occur in the same habi - ever, in the authors’ view, these differences tat are species of the genus Coryphophylax are not sufficiently pronounced to justify the FITZInger in sTeInDAChner , 1867, which recognition of a new genus for the Anda - are primarily inhabiting the understory and manese species. In the absence of a molec - easily differentiated from P. andamanensis ular phylogeny and based on external mor - by the strong transverse skin fold across the phology alone, it is most appropriate to con - shoulder and neck. Calotes cf. versicolor sider this species as a member of the genus (DAuDIn , 1802), probably an introduced Pseudo calotes FITZInger , 1843, which is a species in Andaman Islands, occurs in north highly diverse genus comprising fourteen Andaman and the harbor town of port Blair species and showing a wide array of charac - in south Andaman but is restricted to ters. anthropogenic modified landscapes and not found in forests. Pseudocalotes andama - Pseudocalotes andamanensis nensis is easily distinguished from C. versi - (Boulenger , 1891) color by the regularly arranged, strongly new combination keeled, posterodorsally directed body scales, the relatively short head, and long Chresonyms.- Calotes andamanensis - nuchal and dorsal spines in the latter. Boulenger (1891), sMITh (1935), KrIsh - The herpetofauna of Andaman Islands nAn (2008). is known to have affinities towards the Harikrishnan_Vasudevan_Calotes_andamanensis_HERPETOZOA.qxd 21.06.2013 12:18 Seite 10

12 s. hArIKrIshnAn & K. V AsuDeVAn

Indo-Chinese fauna rather than south Asian In addition, there are several small islands fauna ( sMITh 1940; D As 1999). The chan - within this narrow gap that may have acted nel that separates the Andaman Islands from as stepping-stones for species to colonize the Irrawaddi river Delta of is the Andaman Islands from Myanmar. It is only about 230 km wide at present, but may also likely that increased discharge from the have been as narrow as 70 km during the Irrawaddy river during the monsoons could last glacial maxima ( VorIs 2000; A lFAro et have resulted in a high amount of debris al. 2004). The possibility of a land connec - flowing out into the Andaman sea, that tion between the Andaman Islands and could have acted as rafts for species for col - Myanmar cannot be completely ruled out, onization ( AlFAro et al. 2004). as this is a tectonically, highly active area.

ACKnowleDgMenTs

we would like to thank p. r. sinha (Director, Zoological Museum, university of Copenhagen, for wildlife Institute of India) and V. B. Mathur, (Dean of providing photographs of the holotype of Calotes an- Academics, wildlife Institute of India) for the opportu - damanensis . we also thank the Director of the Zoolo- nity to conduct research at the wildlife Institute of gical survey of India, Dr. B. h. C. Murthy and Dr. India, Dehra Dun. our sincere thanks to the Depart- Kaushik Deuti for facilitating the examination of spec - ment of Forests and wildlife, Andaman and nicobar imens in the collection of the ZsI, Kolkata. we are Islands for permission to conduct fieldwork in the grateful to The Andaman and nicobar environmental Andaman and nicobar Islands. we also thank the Team (AneT) for logistical support. Finally, sincere Department of science and Technology (DsT), gov- thanks are due to s. r. Chandramouli, Anand James ernment of India, for providing monetary support grant Tirkey and sudhir Kumar ekka for help with fieldwork no. sr/so/As-08/2009. we thank Mogens Andersen, in the Andaman Islands.

reFerenCes

AlFAro , M. e. & K Arns , D. r. & V orIs , h. K. IshwAr , n. M. & D As , I. (1998): rediscovery of & A BernAThy , e. & s ellIns , s. l. (2004): phylogeny Calotes andamanensis Boulenger (1891) and a re- of Cerberus (serpentes: homalopsinae) and phylo - assessment of the type locality.- Journal of the Bombay geography of Cerberus rynchops : diversification of a natural history society, Mumbai; 95: 513-514. coastal marine snake in .- Journal of KrIshnAn , s. (2008): new species of Calotes Biogeo graphy, oxford; 31: 1277-1292. (reptilia; squamata: Agamidae) from the southern Boulenger , g. A. (1891): on new or little- western ghats, India.- Journal of herpetology, houston known Indian and Malayan and batrachians.- etc.; 42 (3): 530-535. Annals and Magazine of natural history, london; (6) MAhony , s. (2010): systematic and taxomonic 8: 288-292. revaluation of four little known Asian agamid species, DAs , I. (1999): Biogeography of the amphibians Calotes kingdonwardi sMITh , 1935, Japalura kaul- and reptiles of the Andaman and nicobar Islands, backi sMITh , 1937, Salea kakhienensis AnDerson , India; pp. 43-77. In: oTA , h. (ed.) Tropical island her - 1879 and the monotypic genus Mictopholis sMITh , petofauna. origin, current diversity and current status. 1935 (reptilia: Agamidae).- Zootaxa, Auckland; 2514: Amsterdam (elsevier). 1-23. De roepsTorFF , F. A. (1875): Vocabulary of MAlhoTrA , A. & T horpe , r. s. (1997): new dialects spoken in the nicobar and Andaman Isles, with perspectives on the evolution of south-east Asian a short account of the natives, their customs and habits, pitvipers (genus Trimeresurus ) from molecular studies; and of previous attempts at colonization; Calcutta pp. 115-128. In: Thorpe , r. s. & w üsTer , w. & M Al- (office of the superintendent of government printing), hoTrA , A. (eds.): Venomous snakes: ecology, evolution pp. 140. and snakebite. oxford (The Zoological society of lon- hAllerMAnn , J. (2005): A taxonomic review don/Clarendon press). of the genus Bronchocela (squamata: Agamidae) MooDy , s . M. (1980): phylogenetic and histori - with the description of a new species from .- cal biogeographical relationships of the genera in the russian Journal of herpetology, Moscow; 12 (3): Agamidae (reptilia; lacertilia) . unpublished 167-182. thesis (The university of Michigan), pp. 373. hAllerMAnn , J. & B öhMe , w. (2000): A review sMITh , M . A. (1935): The fauna of British India, of the genus Pseudocalotes (squamata: Agamidae), including Ceylon and Burma - reptilia and Amphibia, with description of a new species from west Malaysia.- Vol. II; london (Taylor & Francis), pp. 440, 1 plate. Amphibia-reptilia, leiden; 21: 193-210. sMITh , M. A. (1940): The herpetology of the hAllerMAnn , J. & M CguIre , J. A. (2001): A Andaman and nicobar Islands.- proceedings of the new species of Pseudocalotes (squamata: Agamidae) linnean society of london, london; 3: 150-158. from Bukit larut, west Malaysia.- herpetologica, sTeInDAChner , F. (1867): reise der österreichi- lawrence; 57 (3): 255-265. schen Fregatte novara um die erde in den Jahren Harikrishnan_Vasudevan_Calotes_andamanensis_HERPETOZOA.qxd 21.06.2013 12:18 Seite 11

rediscovery of Calotes andamanensis Boulenger , 1891 13

1857, 1858, 1859 unter den Befehlen des Commodore western ghats.- Journal of herpetology, lawrence; 45 B. von wüllersTorF -u rBAIr . Zoologischer Theil, (1): 100-110. erster Band. reptilien. wien (gerolds sohn), pp. 98, 3 VorIs , h. K. (2000): Maps of pleistocene sea pls. levels in southeast Asia: shorelines, river systems and Vogel , g. & VAn rooIJen , J. (2011): Contrib- time durations.- Journal of Biogeography, oxford; 27: utions to a review of the Dendrelaphis pictus (gMelIn , 1153-1167. 1789) complex (serpentes: Colubridae) – 3. The Indian wüsTer , w. (1998): The cobras of the genus forms, with the description of a new species from the Naja in India.- hamadryad, Madras 23 (1): 15-32.

AppenDIx : specimens examined

Acronyms: BnhM – Bombay nat- (gunTher , 1870) – ural history society (Mumbai, India); ZsI 6571, yunnan, southern China; Mnhn – Muséum national d’histoire DuMérIl & B IB- naturelle (paris, France); wII – wildlife ron , 1837 – ZsIC 2049, Manipur, India; Institute of India, Vertebrate Collections Calotes versicolor (DAuDIn , 1802) – (Dehra Dun, uttarakhand, India); ZsI – ZsI 3876, 16 specimens from Chipo, north Zoological survey of India, Marine Andaman; ZsI 3877, 2 specimens from regional station (port Blair, Andaman and Chipo, north Andaman; nicobar Islands, India); ZsIC – Zoological Pseudocalotes andamanensis (B ou- survey of India (Kolkata, India). lenger , 1891) – wII 001, long Island, Bronchocela cristatella (Kuhl , 1820) Andaman & nicobar Islands, India; wII – ZsIC 5337, near Tavoy; ZsI 3878- 055, wII 056, wII 072, wII 125, south sK04CnC-8, 11, 16, Car nicobar; Andaman Island, Andaman & nicobar Bronchocela danieli (TIwArI & B Is- Islands, India; wII 126 rutland Island, wAs , 1973) – ZsIC 22455 (holotype), Camp - Andaman & nico bar Islands, India; bell Bay, great nicobar, Andaman & nico - Pseudocalotes austeniana (AnnAn- bar Islands, India; ZsI 3879-sK03ln-34, DAle , 1908) – (holotype), ZsI 3976, hills little nico bar; near harmatti, Assam; Bronchocela rubrigularis hAller - Pseudocalotes microlepis (B oulen - MAnn , 2009 – ZsI 3880-sK03nC1, nan - ger , 1888) – ZsIC 11940, Manipur, India; cowry; Pseudocalotes tympanistriga (grAy , Calotes aurantolabium KrIshnAn , 1831) – Mnhn 6889, 6669, 11-147-148 and 2008 – BnhM 1436, (holotype), Kalakkad 95-475. Mundanthurai Tiger reserve, Tamil nadu, India;

DATe oF suBMIssIon: July 13, 2012 Corresponding editor: heinz grillitsch

AuThors: surendran hArIKrIshnAn , Karthikeyan VAsuDeVAn (corresponding author < karthik@wii. gov.in >), wildlife Institute of India, Chandrabani, Dehradun 248001, uttarakhand, India